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Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39. With 7 figures<br />

<strong>The</strong> <strong>orchid</strong> <strong>flora</strong> <strong>of</strong> <strong>Cocos</strong> <strong>Island</strong> <strong>National</strong> <strong>Park</strong>,<br />

Puntarenas, Costa Rica<br />

DIEGO BOGARÍN 1 *, JORGE WARNER 1 , MARTYN POWELL 2 and<br />

VINCENT SAVOLAINEN 2,3<br />

1 Jardín Botánico Lankester, Universidad de Costa Rica. P.O. Box 302-7050 Cartago, Costa Rica, A.C.<br />

2 Imperial College London, Silwood <strong>Park</strong>, Ascot, Berkshire, SL5 7PY, UK<br />

3 Royal Botanic Gardens, Kew, Richmond, Middlesex, TW9 3DS, UK<br />

Received 13 January 2011; revised 31 January 2011; accepted for publication 2 February 2011<br />

<strong>The</strong> <strong>orchid</strong> <strong>flora</strong> <strong>of</strong> <strong>Cocos</strong> <strong>Island</strong> <strong>National</strong> <strong>Park</strong>, Costa Rica is described based on field collections and herbarium<br />

sampling. <strong>Cocos</strong> <strong>Island</strong> is an oceanic island situated 550 km south-west <strong>of</strong> Puntarenas (the main Costa Rican port<br />

on the Pacific coast) and 680 km north-east <strong>of</strong> the Galápagos <strong>Island</strong>s. Five species <strong>of</strong> <strong>orchid</strong>s were recorded in this<br />

small area <strong>of</strong> 24 km 2 : Camaridium micranthum and Ornithidium adendrobium (both formerly included in<br />

Maxillaria) and three endemic species <strong>of</strong> Epidendrum (E. cocoense, E. insulanum and E. jimenezii). <strong>The</strong> species are<br />

described and analytical illustrations are provided for each. A key to the species for field identification based on<br />

morphology is presented. Biogeography, ecology, taxonomy, evolution and conservation status <strong>of</strong> the species are also<br />

discussed. © 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39.<br />

ADDITIONAL KEYWORDS: Camaridium – Epidendrum – floristics – Maxillaria – oceanic islands –<br />

Orchidaceae – Ornithidium – protected areas – taxonomy.<br />

INTRODUCTION<br />

Isla del Coco, also known as <strong>Cocos</strong> <strong>Island</strong>, is an<br />

oceanic island in the south-eastern tropical Pacific<br />

Ocean (5°31′39″N, 87°03′32″W), situated 550 km<br />

south-west <strong>of</strong> Puntarenas, the main port <strong>of</strong> Costa Rica<br />

on the Pacific coast (Fig. 1). <strong>The</strong> nearest continental<br />

point is Cabo Blanco on the Península de Nicoya,<br />

Costa Rica, 532 km from the island. Other oceanic<br />

islands relatively near <strong>Cocos</strong> <strong>Island</strong> are Isla Pinta <strong>of</strong><br />

the Galápagos archipelago (Ecuador) at 681 km, Isla<br />

Malpelo (Colombia) at 630 km, Isla Coiba (Panama)<br />

at 632 km and Clipperton <strong>Island</strong> (Île de Clipperton or<br />

Île de la Passion, France) at 2375 km.<br />

<strong>Cocos</strong> <strong>Island</strong> became part <strong>of</strong> Costa Rica in 1869,<br />

when President Jesús Jiménez decreed it as national<br />

territory. At that time, the island was uninhabited and<br />

had never been claimed by any other country. Its name<br />

was probably a mistranscription from the original map<br />

*Corresponding author. E-mail: diego.bogarin@ucr.ac.cr<br />

20<br />

by the French cartographer Nicolas Desliens in 1541,<br />

in which the island appeared (for first time) as ‘Y e<br />

Coques’ (Hogue & Miller, 1981; Trusty, 2004). In<br />

French, the word ‘coque’ means eggshell or nutshell<br />

and the name could have been a reference to the<br />

common fruit <strong>of</strong> the abundant endemic tree Sacoglottis<br />

holdridgei Cuatrec. (Humiriaceae).<br />

In 1978, under the government <strong>of</strong> President<br />

Rodrigo Carazo, the island was decreed a <strong>National</strong><br />

<strong>Park</strong> and a zone <strong>of</strong> absolute protection. Administratively,<br />

the area belongs to the Area de Conservación<br />

Marina Isla del Coco (ACMIC) <strong>of</strong> the Costa Rican<br />

Ministry <strong>of</strong> Environment, Energy and Telecommunications<br />

(MINAET) and its <strong>National</strong> System <strong>of</strong> Conservation<br />

Areas (SINAC). Later, in 1997 the island<br />

became a UNESCO-designated World Natural Heritage<br />

Site in recognition <strong>of</strong> its exceptional biodiversity.<br />

In 2002, the World Natural Heritage Site designation<br />

was extended to include an expanded marine zone<br />

<strong>of</strong> 1997 km 2 , thus protecting marine ecosystems,<br />

populations <strong>of</strong> sharks, rays, dolphins and other large<br />

marine species. In addition, in 1998 the island was<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39<br />

ORCHID FLORA OF COCOS ISLAND 21<br />

Figure 1. Map <strong>of</strong> <strong>Cocos</strong> <strong>Island</strong> showing collecting sites. Scale 1 : 25 000.


22 D. BOGARÍN ET AL.<br />

included in the list <strong>of</strong> Wetlands <strong>of</strong> International<br />

Importance under the RAMSAR Convention.<br />

TOPOGRAPHY<br />

<strong>The</strong> total land area is 24.85 km2 (c. 7.6 ¥ 4.4 km) with<br />

the following limits: to the north Punta Agujas, to the<br />

south Cabo Dampier, to the west Cabo Lionel and to<br />

the east Cabo Atrevido (Fig. 1). <strong>The</strong> topography is<br />

abrupt with high slopes and cliffs. <strong>The</strong> highest point<br />

is Cerro Iglesias, observable as a cloudy mountain<br />

rising 575 m above sea level. Other important hills<br />

are Cerro Pelón at 530 m and Cerro Jesús Jiménez at<br />

430 m <strong>of</strong> elevation. Some flat areas are observed in<br />

the region called Llanos Palo de Hierro at 300 m<br />

elevation (Fig. 1). <strong>The</strong> island has four bays: Iglesias,<br />

Weston, Chatham and Wafer. <strong>The</strong> later two are the<br />

most important for boats and crafts and have the<br />

main ranger stations. <strong>The</strong> most important rivers are:<br />

the Genio to the north-east that flows into Wafer Bay,<br />

the Chatham River that flows into Chatham Bay and<br />

the Iglesias to the south, which terminates in Iglesias<br />

Bay. <strong>The</strong> irregular topography produces many waterfalls<br />

because <strong>of</strong> the vertical cliffs and high water<br />

availability in the soil.<br />

GEOLOGY<br />

<strong>Cocos</strong> <strong>Island</strong> emerged from the ocean through volcanic<br />

activity <strong>of</strong> the submarine mountain range called<br />

the <strong>Cocos</strong> Ridge. <strong>The</strong> ridge was formed in the Miocene<br />

by tectonic movements <strong>of</strong> the <strong>Cocos</strong> and Nazca plates<br />

and volcanic action <strong>of</strong> the Galápagos hotspot. <strong>The</strong>se<br />

geological events occurred along the Pacific Ocean<br />

near South America. <strong>The</strong>n, the island moved to the<br />

north as a result <strong>of</strong> the action <strong>of</strong> the <strong>Cocos</strong> and Nazca<br />

plates. Geologists have estimated the age <strong>of</strong> the<br />

island to be 1.9–2.4 million years (Bellon, Sáenz &<br />

Tournon, 1984). <strong>The</strong> soils are composed <strong>of</strong> basaltic<br />

and pyroclastic alkaline rocks, the remains <strong>of</strong> ancient<br />

volcanic activity. However, the acidic condition is<br />

caused by the degradation <strong>of</strong> organic material under<br />

high humidity and the consequent action <strong>of</strong> bacteria<br />

and fungi (Trusty, 2004).<br />

CLIMATE<br />

<strong>The</strong> island is under the influence <strong>of</strong> the Intertropical<br />

Convergence Zone (ITCZ) and receives other climatic<br />

influences such as the El Niño-Southern Oscillation<br />

(ENSO). <strong>The</strong> climate <strong>of</strong> the island was discussed by<br />

Alfaro (2008) and Lizano (2008). A complex system <strong>of</strong><br />

marine currents originating in America and the Indo-<br />

Pacific region flows around the island, thus influencing<br />

its climate (Lizano, 2008). <strong>The</strong> island experiences<br />

high rainfall, which can reach 7000 mm per year.<br />

<strong>Cocos</strong> is the only island in the tropical eastern Pacific<br />

with a tropical humid forest. According to Tosi (1969),<br />

the life zone is the premontane belt forest, basal belt<br />

transition. <strong>The</strong> period <strong>of</strong> high rainfall is between May<br />

and October. <strong>The</strong>re is no true dry season but rainfall<br />

is slightly lower from November to March (Alfaro,<br />

2008). <strong>The</strong> average temperature is approximately<br />

27–28 °C. <strong>The</strong>se climatic factors allow the establishment<br />

<strong>of</strong> plants such as bromeliads, ferns and epiphytic<br />

<strong>orchid</strong>s.<br />

BIODIVERSITY<br />

<strong>The</strong> biodiversity <strong>of</strong> <strong>Cocos</strong> <strong>Island</strong> includes both marine<br />

and terrestrial ecosystems. Marine biodiversity is the<br />

main attraction for touristic and scientific activities.<br />

This place is an important breeding site for many<br />

species. <strong>The</strong> migration and reproduction <strong>of</strong> sharks is<br />

one <strong>of</strong> the most spectacular marine attractions for<br />

divers. It is possible to observe the rare and endangered<br />

whale shark (Rhincodon typus) and large populations<br />

<strong>of</strong> hammerhead sharks (Sphyrna lewinni), the<br />

marine emblem <strong>of</strong> the island. <strong>The</strong> history <strong>of</strong> marine<br />

research at <strong>Cocos</strong> <strong>Island</strong> was reviewed recently by<br />

Cortés (2008). Terrestrial biodiversity is also important.<br />

According to the most recent botanical treatment<br />

<strong>of</strong> the island by Trusty, Kesler & Haug Delgado<br />

(2006), there are 262 species <strong>of</strong> plants, <strong>of</strong> which 37<br />

(approximately 14%) are endemic. Among the terrestrial<br />

fauna, there are 382 species <strong>of</strong> insects, five<br />

species <strong>of</strong> reptiles, one scorpion and 12 species <strong>of</strong><br />

birds, including the endemic Coco finch, Pinaroloxias<br />

inornata. <strong>The</strong>re is a low elevation rainforest formed<br />

mainly by the endemic tree Sacoglottis holdridgei. At<br />

the highest points (i.e. Cerro Iglesias), there is a high<br />

elevation cloud forest. Other important plant communities<br />

are the riparian habitats, estuarine habitats,<br />

coastal cliff communities, shore vegetation, landslides<br />

and rocky islets that have sparse grassy vegetation<br />

and scattered trees <strong>of</strong> Clusia rosea Jacq. <strong>The</strong>se plant<br />

associations were defined and characterized by Trusty<br />

(2004). Detailed information on climate, geology,<br />

human impacts and introduced animals and plants<br />

can be found in Trusty (2004) and Trusty et al. (2006).<br />

THE HISTORY OF COCOS ISLAND ORCHIDS<br />

<strong>The</strong> first recorded visit to <strong>Cocos</strong> <strong>Island</strong> was in 1526,<br />

when the Spanish explorer Joan Cabeças reached the<br />

island. Since then, mariners and pirates have known<br />

the island and it became popular because <strong>of</strong> traditional<br />

stories <strong>of</strong> treasure supposedly buried there.<br />

According to the legend, two treasures have been<br />

hidden on <strong>Cocos</strong> <strong>Island</strong>: firstly, the pirate Bennett<br />

Graham (Benito Bonito) <strong>of</strong> the Relámpago, who looted<br />

several cities around South America, is reputed to<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


have kept his treasure in an unknown place on the<br />

island (Hogue & Miller, 1981; Trusty, 2004) and, secondly,<br />

the ‘treasure <strong>of</strong> Lima’ <strong>of</strong> Captain Thompson <strong>of</strong><br />

the Mary Dear may be on <strong>Cocos</strong>, including priceless<br />

gold and jewels from Peru. Following these tales,<br />

people looking for treasures made > 400 expeditions<br />

to the island and <strong>Cocos</strong> is therefore also well known<br />

as the ‘treasure island’.<br />

In spite <strong>of</strong> these hundreds <strong>of</strong> expeditions, plant<br />

collections only started in 1836 when the English<br />

collector George W. Barclay visited <strong>Cocos</strong> <strong>Island</strong> as<br />

part <strong>of</strong> an exploration <strong>of</strong> the Pacific coast aboard<br />

H.M.S. Sulphur, commanded by Captain Sir Edward<br />

Belcher (Bentham, 1844–1846). However, in their<br />

botanical treatment they did not report or describe<br />

any <strong>orchid</strong> species. After Barclay’s expedition, the<br />

marine zoologist and oceanographer Alexander<br />

Agassiz collected plants during the late 19 th century<br />

(1888 and 1891). Aboard the Albatross, he visited<br />

<strong>Cocos</strong> <strong>Island</strong> on his way to the Galápagos. He collected<br />

Epidendrum insulanum Schltr. (then undescribed),<br />

the first <strong>orchid</strong> specimen collected in the<br />

island (Agassiz s.n., GH).<br />

In 1898, the government <strong>of</strong> President Rafael Yglesias<br />

sponsored the first Costa Rican expedition. He<br />

assigned to the scientists Anastasio Alfaro and Henry<br />

Pittier the task <strong>of</strong> assessing the island in order for<br />

it to be reopened as a penal center (Pittier, 1898).<br />

<strong>The</strong>y went aboard the Poás and collected plants<br />

and animals. Four years later, on a second trip in<br />

1902, together with Paul Biolley, aboard the Turrialba,<br />

Pittier collected two specimens <strong>of</strong> Epidendrum<br />

L. One was the first <strong>orchid</strong> species described from the<br />

island, published by R. Schlechter as E. insulanum<br />

in Beihefte zum Botanischen Centralblatt in 1918.<br />

Unfortunately, the specimen studied by Schlechter<br />

was destroyed in the Dahlem-Berlin Herbarium<br />

during the Second World War and there is no indication<br />

that Schlechter received more than one specimen<br />

from Pittier. However, Pittier prepared at least five<br />

duplicates; two are kept at the herbarium <strong>of</strong> the<br />

Museo Nacional de Costa Rica (CR) and three at<br />

Oakes Ames Orchid Herbarium <strong>of</strong> Harvard University<br />

(AMES), GH-3581 (=AMES-70445), GH-3579<br />

(=AMES-70446) and the lectotype designated by<br />

Trusty & Blanco (2005) GH-3580 (=AMES-73449).<br />

<strong>The</strong>se specimens are available under Pittier n. 16350<br />

(Institute. Phys. Geogr. Cost.); the same number is<br />

cited by Schlechter (1918) in the protologue. A sheet<br />

with tracings <strong>of</strong> Schlechter’s drawing <strong>of</strong> the holotype<br />

is also kept at AMES (AMES-70447). <strong>The</strong> second<br />

species recorded by Pittier correspond to at least two<br />

plants <strong>of</strong> E. cocoense Hágsater (then undescribed)<br />

kept at AMES under Pittier n. 16351 (Institute. Phys.<br />

Geogr. Cost.). Hágsater (1999a) identified the specimens<br />

as E. cocoense. As Schlechter did not mention<br />

ORCHID FLORA OF COCOS ISLAND 23<br />

any <strong>of</strong> these specimens or describe another species, it<br />

is probable that Pittier did not send any more specimens<br />

to Berlin, but no information is available to<br />

clarify this situation.<br />

Robert E. Snodgrass and Edmund Heller in the<br />

Hopkins-Stanford Expedition made another expedition<br />

in 1899. Alban Stewart collected again for the<br />

California Academy <strong>of</strong> Sciences in 1905. <strong>The</strong>n, Henry<br />

Knute Svenson, who was the curator <strong>of</strong> Brooklyn<br />

Botanical Garden, New York, spent 3 days collecting<br />

plants in the island as the chief botanist on the<br />

Vincent Astor Expedition in April 1930 (Svenson,<br />

1935). Svenson found what is now the type specimen<br />

<strong>of</strong> E. cocoense growing by a stream flowing into Wafer<br />

Bay (H. K. Svenson 333, AMES-37853); the species<br />

was later described by Hágsater (1999a). Svenson<br />

prepared two duplicates; one is at AMES and the<br />

other at Chicago (F).<br />

Svenson was the first to collect specimens <strong>of</strong> Camaridium<br />

Lindl. (the specimens from the island were<br />

formerly included in Maxillaria Ruiz & Pav.). <strong>The</strong>y<br />

were misidentified under the names M. concavilabia<br />

Ames & Correll and Ornithidium aff. stenophyllum<br />

Schltr. In 1994, John T. Atwood identified the<br />

specimens as M. parvi<strong>flora</strong> (Poepp. & Endl.)<br />

Garay (=Camaridium micranthum M.A.Blanco, see<br />

Discussion).<br />

Two years after Svenson’s trip, the American botanist<br />

John Thomas Howell explored the island on <strong>The</strong><br />

Templeton Crocker Expedition sponsored by the California<br />

Academy <strong>of</strong> Sciences. He collected another<br />

specimen <strong>of</strong> E. cocoense erroneously identified as E.<br />

ramosum Jacq. var. imbricatum Ames, F.T.Hubb. &<br />

C.Schweinf. Later, President Cruise <strong>of</strong> the USA made<br />

other expeditions from 1935 to 1940.<br />

<strong>The</strong> Costa Rican botanist and ex-director <strong>of</strong> the<br />

Museo Nacional de Costa Rica, Juvenal Valerio<br />

Rodríguez, went to the island on two expeditions<br />

during 1939 and 1940. He collected C. micranthum<br />

(misidentified as Maxillaria variabilis Bateman ex<br />

Lindl) and another specimen <strong>of</strong> E. cocoense was collected<br />

in his second trip.<br />

Witold L. Klawe, a fishery researcher, collected<br />

plants as part <strong>of</strong> the Inter-American Tropical Tuna<br />

Commission aboard the Costa Rica Dome Cruise in<br />

1959. He collected E. cocoense, wrongly identified<br />

under the names E. paranaense Barb. Rodr. and E.<br />

santaclarense Ames.<br />

Hágsater (1999b) described the last species known<br />

from the island under the name E. jimenezii. <strong>The</strong><br />

species honours Alfonso Jiménez, ex-director <strong>of</strong> the<br />

Museo Nacional de Costa Rica, who collected the type<br />

specimen in 1965 (A. Jiménez 3178, CR). He stated<br />

that the plant had white flowers and it is apparently<br />

a common epiphyte along the Genio River, near its<br />

mouth into Wafer Bay.<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


24 D. BOGARÍN ET AL.<br />

In 1967, the botanists Ira L. Wiggins and Duncan<br />

M. Porter, who worked on the Flora <strong>of</strong> the Galápagos<br />

<strong>Island</strong>s, collected specimens <strong>of</strong> E. insulanum (Hágsater,<br />

2007). Luis D. Gómez went to the island in 1970<br />

and made another collection <strong>of</strong> E. cocoense from the<br />

forest <strong>of</strong> Wafer Bay.<br />

After many expeditions, the number <strong>of</strong> species <strong>of</strong><br />

the island remained unclear. In 1973, in a trip organized<br />

by the Smithsonian Tropical Research Institute,<br />

Robert L. Dressler collected an Epidendrum, identified<br />

by Hágsater as E. jimenezii (Dressler 4468, PMA-<br />

36761), but a duplicate kept at CR (Dressler 4468,<br />

CR) could be an E. insulanum as noted by Trusty &<br />

Blanco (2006). A mixed collection was probably made<br />

under the same number, demonstrating the difficulties<br />

in identifying those species without flowers as<br />

their vegetative features are highly variable. Another<br />

sample <strong>of</strong> E. cocoense was collected and a new record,<br />

Ornithidium adendrobium (Rchb.f.) M.A.Blanco &<br />

Ojeda (formerly known as Maxillaria adendrobium<br />

(Rchb.f.) Dressler) was found near Wafer Bay. This<br />

widespread species ranges in both continental and<br />

insular lands around the Neotropics. In 1865, Heinrich<br />

G. Reichenbach described it as Ponera adendrobium<br />

Rchb.f. based on a collection <strong>of</strong> Charles Wright<br />

in Cuba.<br />

In 1979, Robin Foster collected an interesting Epidendrum<br />

in the floodplain <strong>of</strong> Genio River. It seems to<br />

be an example <strong>of</strong> E. insulanum, but Hágsater cited<br />

it as E. jimenezii (Hágsater, 1999b). In 1980, Pablo<br />

Sánchez-Vindas collected E. cocoense and in 1981<br />

Jorge Gómez-Laurito collected E. jimenezii (wrongly<br />

identified as Epidendrum imbricatum Lam.) and<br />

another E. insulanum. He also collected E. cocoense,<br />

and Luis D. Gómez collected M. parvi<strong>flora</strong> in the<br />

same year. In 1989, R. Soto collected E. cocoense, O.<br />

adendrobium and other two specimens <strong>of</strong> C. micranthum.<br />

Other collections were made in 1994 by the<br />

researchers <strong>of</strong> Instituto Nacional de Biodiversidad<br />

(INBio). However, they reported the same five<br />

species. With the description <strong>of</strong> E. cocoense and<br />

E. jimenezii by Hágsater (1999b), the <strong>orchid</strong> <strong>flora</strong> <strong>of</strong><br />

<strong>Cocos</strong> <strong>Island</strong> is made up <strong>of</strong> three endemic species <strong>of</strong><br />

Epidendrum (E. cocoense, E. insulanum and E. jimenezii),<br />

one species <strong>of</strong> Camaridium, C. micranthum,<br />

and one species <strong>of</strong> Ornithidium, O. adendrobium<br />

(Dressler, 2003). <strong>The</strong> two former species are also<br />

found in continental parts <strong>of</strong> the Neotropics.<br />

Recently, Trusty and co-workers (Trusty, 2004;<br />

Trusty & Blanco, 2005, 2006; Trusty et al., 2006)<br />

finished their floristic treatment <strong>of</strong> the <strong>Cocos</strong> <strong>Island</strong><br />

<strong>National</strong> <strong>Park</strong>. <strong>The</strong>y reported the same five species<br />

for the island. However, they were unable to locate<br />

material <strong>of</strong> E. jimenezii, and consequently they could<br />

not distinguish this species from E. insulanum<br />

(Trusty & Blanco, 2006).<br />

<strong>The</strong> aim <strong>of</strong> this paper is to present detailed information<br />

for identifying the <strong>orchid</strong>s <strong>of</strong> <strong>Cocos</strong> <strong>Island</strong>, to<br />

provide illustrations <strong>of</strong> the five species based on living<br />

plants (two <strong>of</strong> them are illustrated for a first time)<br />

and to discuss the identity <strong>of</strong> the rare E. jimenezii.<br />

This work is part <strong>of</strong> a series <strong>of</strong> floristic studies<br />

intended to clarify the diversity and taxonomy <strong>of</strong><br />

Orchidaceae in Costa Rica (e.g. Pupulin, 2010) and its<br />

national parks and protected areas (Pupulin, 1998;<br />

Bogarín & Pupulin, 2007).<br />

MATERIAL AND METHODS<br />

This work was conducted mainly in the <strong>Cocos</strong> <strong>Island</strong><br />

<strong>National</strong> <strong>Park</strong>, Puntarenas, Costa Rica and the<br />

Lankester Botanical Garden (LBG), University <strong>of</strong><br />

Costa Rica. Living specimens were collected, cultivated<br />

and documented between 2006 and 2009. Data<br />

from all specimens cited have been recorded in a<br />

computerized database at LBG. <strong>The</strong>y are also available<br />

on the website at http://www.epidendra.org<br />

(Pupulin, 2007; Pupulin, 2009). Georeferences for<br />

specimens were obtained using a Garmin eTrex Vista<br />

GPS and maps. Ecological zones were estimated by<br />

using the Holdridge Life Zone System (Holdridge,<br />

1967, 1987) and the ecological map <strong>of</strong> Costa Rica<br />

(Tosi, 1969).<br />

Phenology data were recorded both in the field and<br />

in cultivated specimens or herbarium labels. Individual<br />

plants were photographed, illustrated and<br />

preserved as herbarium specimens and preserved<br />

specimens in formaldehyde : acetic acid : ethanol<br />

[FAA (53% ethanol, 37% water, 5% formaldehyde and<br />

5% glycerol)] (including flowers, portions <strong>of</strong> the stems<br />

or entire plants) for future reference. Newly collected<br />

herbarium specimens were deposited at CR, JBL and<br />

USJ herbaria. Whenever possible, the herbarium<br />

specimens were complemented with sketches, photographs<br />

and FAA material. <strong>The</strong> material preserved in<br />

FAA was deposited at JBL and indicated in the treatment<br />

as ‘JBL-spirit’. Herbarium and spirit material<br />

may consist <strong>of</strong> wild collected specimens or material<br />

collected entirely from cultivated plants.<br />

Sketches <strong>of</strong> specimens were made with a Leica<br />

MZ7.5 stereomicroscope with drawing tube and conserved<br />

in the reference collections at JBL. All taxa<br />

were illustrated by composite line drawings from<br />

living specimens. Illustrations include a typical plant<br />

habit, inflorescences or part <strong>of</strong> the inflorescences, the<br />

flower and a dissection <strong>of</strong> perianth. Plate composition<br />

was as consistent as possible to facilitate species<br />

comparison.<br />

Descriptions were prepared from both living specimens<br />

and herbarium material. Materials from the<br />

following herbaria were studied: AMES, BM, CAS,<br />

CR, INB, JBL, K, PMA and USJ.<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


KEY TO THE ORCHIDS OF THE COCOS ISLAND NATIONAL PARK<br />

LIST OF SPECIES<br />

1. CAMARIDIUM LINDL.<br />

CAMARIDIUM LINDL., Bot. Reg. 10: sub t. 844. 1824.<br />

Type species: Camaridium ochroleucum Lindl., Bot.<br />

Reg. 10: 844. 1824.<br />

PLANT epiphytic, caespitose or straggling plants,<br />

small to large in size, variable in habit. PSEUDOBULBS<br />

unifoliate, separated by an elongate rhizome, caespitose<br />

or lacking pseudobulbs with erect, monopodial<br />

leafy shoots. Rarely with dimorphic growth. LEAVES<br />

conduplicate, thin or coriaceous to subcoriaceous, conduplicate.<br />

INFLORESCENSES produced from a leafy<br />

new growth. From several to many. FLORAL BRACT<br />

longer than the ovary. FLOWERS small and inconspicuous<br />

to medium sized and showy. SEPALS AND<br />

PETALS subsimilar, spreading. LIP <strong>of</strong>ten shorter than<br />

the other perianth parts and <strong>flora</strong>l segments, lacking<br />

fibres, simple or three-lobed, articulated with or<br />

adnate to the column, usually provided at the base<br />

with a fleshy callus. COLUMN erect, terete. CAPSULES<br />

have apical dehiscence.<br />

Camaridium is a Neotropical genus with wide distribution<br />

and approximately 80 species. <strong>The</strong> highest<br />

diversity is found in Central America.<br />

1. CAMARIDIUM MICRANTHUM M.A.BLANCO, Lankesteriana<br />

7: 520. 2007. Scaphyglottis parvi<strong>flora</strong> Poepp.<br />

& Endl., nov. gen. ac Sp. 1: 58, t. 97. 1835. Maxillaria<br />

parvi<strong>flora</strong> (Poepp. & Endl.) Garay, Bot. Mus. Leafl.<br />

21: 258. 1967. Non Camaridium parviflorum Fawc.,<br />

Symb. Antill. 1: 472. 1910. Type: PERU. Crescit ad<br />

arbores Peruviae in viciniis praedii Cuchero, floret<br />

Feb, Poeppig s.n. (holotype: W). (Figs 2, 3A, B).<br />

Description: PLANT epiphytic, pendent or suberect, to<br />

25 cm long. ROOTS flexuous, < 1.5 mm in diameter,<br />

white to greenish, with green or orange–yellowish<br />

tips. PSEUDOBULBS unifoliate, ovoid, pyriform, separated<br />

by an elongate rhizome, 2.5–6 ¥ 0.8–1.3 cm,<br />

internodes up to 3 cm long, covered by scarious,<br />

imbricate papyraceous sheaths. LEAVES apical on<br />

ORCHID FLORA OF COCOS ISLAND 25<br />

1. Plants with pseudobulbs, each with one apical leaf.................................................Camaridium micranthum<br />

1. Plant without pseudobulbs, the leaves distichously arranged.....................................................................2<br />

2. Inflorescense lateral, lip adnate to the column foot................................................Ornithidium adendrobium<br />

2. Inflorescence apical, lip adnate to the footless column..............................................................................3<br />

3. Inflorescence with one green–yellow flower < 1 cm in length, apex <strong>of</strong> column erose, lateral lobes <strong>of</strong> lip not covering<br />

the column apex .................................................................................................Epidendrum insulanum<br />

3. Inflorescence with two or more white to whitish flowers > 1 cm in length, column apex with two teeth in each side,<br />

lateral lobes <strong>of</strong> lip covering the column apex..........................................................................................4<br />

4. Leaves 1.5–2.7 cm wide, <strong>flora</strong>l bracts 1.3–1.6 cm wide, broadly ovate, larger than the scurfy ovary...Epidendrum<br />

cocoense<br />

4. Leaves 1.0–1.2 cm wide, <strong>flora</strong>l bracts 0.5–0.6 cm wide, tubular, as long as the glabrescent ovary......Epidendrum<br />

jimenezii<br />

each pseudobulb, petiolate, oblong–linear, acute,<br />

emarginate, unequally two-lobed at the apex, conduplicate,<br />

subcoriaceous, to approximately 10–20 ¥ 1.0–<br />

1.4 cm. INFLORESCENCE a single-flowered raceme,<br />

produced in clusters along the stem, at the base <strong>of</strong><br />

each pseudobulb, covered by several scarious, imbricate,<br />

acute, papyraceous sheaths. PEDICEL inconspicuous.<br />

FLORAL BRACTS triangular, ovate, scarious.<br />

OVARY cylindrical, < 4 mm long. FLOWERS small,<br />

inconspicuous, white with a yellow dot in the lip<br />

callus, approximately 5 mm in length. DORSAL SEPAL<br />

oblong, acute, conduplicate, concave towards the<br />

apex, 5 ¥ 2 mm. LATERAL SEPALS narrowly ovate,<br />

acute, slightly conduplicate, concave, 4.5 ¥ 2.5 mm.<br />

PETALS narrowly oblong, acute, 3.8 ¥ 0.8 mm. LIP<br />

simple, spathulate, oblong, obtuse, attached to the<br />

column foot, conduplicate, callus laminate, 3.0 ¥<br />

1.5 mm. COLUMN short, terete, to 1 mm long, anther<br />

apical, stigma ventral, rectangular. POLLINIA four,<br />

ovoid, with a short stipe, viscidium elliptic. ANTHER<br />

CAP subquadrate–cucullate.<br />

Other vouchers examined: COSTA RICA. Puntarenas:<br />

Puntarenas, Isla del Coco, cumbre del Cerro Iglesias,<br />

575 m, 5°31′45.8N, 87°04′80.5″W, bosque pluvial premontano,<br />

transición a basal, epífitas en Sacoglottis<br />

holdridgei, 15 Abril 2006, D. Bogarín 2766, G. Blanco,<br />

G. Gigot, V. Savolainen & J. Warner (JBL-Spirit);<br />

Puntarenas: Puntarenas, Isla del Coco, camino entre<br />

Bahía Wafer y Bahía Chatham, orillas del sendero,<br />

210 m, 5°32′84.07″, N 87°03′0.18″W, bosque pluvial<br />

premontano, transición a basal, epífitas en Sacoglottis<br />

holdridgei, 14 abril 2006, D. Bogarín 2766, G. Blanco,<br />

G.Gigot, V. Savolainen & J. Warner (JBL-Spirit);<br />

Puntarenas: Puntarenas, Isla del Coco, camino al<br />

Cerro Iglesias, ascenso en el primer mirador, 275 m,<br />

5°32′14.09N, 87°03′36.8″W, bosque pluvial premontano,<br />

transición a basal, epífitas en Sacoglottis holdridgei,<br />

15 abril 2006, D. Bogarín 2746, G. Blanco,<br />

G.Gigot, V. Savolainen & J. Warner (JBL-Spirit). Pun-<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


26 D. BOGARÍN ET AL.<br />

Figure 2. Camaridium micranthum M.A.Blanco. A, habit. B, flower. C, dissected perianth. D, column and lip, lateral<br />

view. E, column, lateral and front views. F, pollinarium anther cap. Drawn by D. Bogarín from Bogarín 2746 (JBL-Spirit).<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


ORCHID FLORA OF COCOS ISLAND 27<br />

Figure 3. Floral morphology <strong>of</strong>: (A, B) Camaridium micranthum; (C) Epidendrum cocoense; (D) Epidendrum insulanum;<br />

(E) Epidendrum jimenezii; (F) Ornithidium adendrobium.<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


28 D. BOGARÍN ET AL.<br />

tarenas: Isla del Coco. Bahía Chatham, 5°32′82″N,<br />

82°02′17″W, c. 200 m, 12 marzo 1994, G. Dauphin<br />

1175 (CR, USJ); J. Gómez 18043 (CR); Quesada 1081<br />

(INB); A. Rojas 3659 (CR, INB); 6–8 junio 1989, R.<br />

Soto s.n. (USJ-49289 and 49290); along a brook<br />

flowing into Wafer Bay, April 18, 1930, H. K. Svenson<br />

334 (GH); <strong>Cocos</strong> <strong>Island</strong>. Closed canopy forest, 5°31′N,<br />

87°03′W, 26 January 2002, J. Trusty 104, 156, 244,<br />

482, and 504 (CR); Isla del Coco, diciembre 1939,<br />

J. Valerio 1086 and 1106 (CR).<br />

Distribution: this species ranges from South Florida,<br />

Antilles and Mexico to Honduras and from Colombia<br />

to Brazil. It seems oddly absent from Nicaragua to<br />

Panama. In Costa Rica it is known only from <strong>Cocos</strong><br />

<strong>Island</strong>.<br />

Etymology: From the Greek micranthum, ‘small<br />

flower’, in allusion to the small and inconspicuous<br />

flowers <strong>of</strong> this species.<br />

Phenology: September to April<br />

Habitat and ecology: a common epiphyte in premontane<br />

rainforest, basal belt transition. It is found from<br />

sea level to the top <strong>of</strong> Cerro Iglesias at 585 m. <strong>The</strong>y<br />

flower throughout the year but mostly between<br />

November to April Pollination is possibly carried out<br />

by ants (Singer, 2003). <strong>The</strong>re are no bees in the island<br />

but there are several species <strong>of</strong> ants. Apparently, the<br />

plants are not self-pollinated. Some plants in cultivation<br />

at Lankester Botanical Garden did not produce<br />

fruits.<br />

Discussion: This is the only species on the island<br />

having pseudobulbs with one apical leaf and the<br />

pseudobulbs are produced in chains. It can be recognized<br />

by the inconspicuous flowers in clusters at the<br />

base <strong>of</strong> the pseudobulbs, the partially connate lateral<br />

sepals, the nearly absent mentum and the cuneate<br />

lip. Poeppig and Endlicher described this species in<br />

Nova Genera ac Species Plantarum under the name<br />

Scaphyglottis parvi<strong>flora</strong> in 1835. <strong>The</strong> name was based<br />

on a plant collected in Cuchero, Peru. Garay transferred<br />

this species to Maxillaria (Garay, 1967).<br />

Molecular studies and nomenclatural changes in<br />

Maxillariinae Benth. & Hook.f. (Blanco et al., 2007;<br />

Whitten et al., 2009) revealed that this species should<br />

be placed under the resurrected Camaridium. Asthe<br />

name Camaridium parviflorum Fawc. was previously<br />

applied to a different species (a synonym <strong>of</strong> O. adendrobium,<br />

a species also occurring in the island), thus<br />

preventing its use in Camaridium, a new name, C.<br />

micranthum, was proposed by Blanco et al. (2007).<br />

Conservation status: This species is not rare and has<br />

a wide distribution range throughout Neotropics.<br />

In the island it is a common epiphyte. This species<br />

should not be regarded as endangered and may well<br />

fit the IUCN Least Concern (LC) category for widespread<br />

and abundant taxa.<br />

2. EPIDENDRUM L.<br />

EPIDENDRUM L., Sp. Pl. ed. 2: 1347. 1763., nom. cons.<br />

Type species: Epidendrum nocturnum Jacq., type.<br />

cons., Enum. Pl. Carib. 29. Jan–December 1760.<br />

PLANT epiphytic or rarely terrestrial herbs to subshrubs,<br />

small to large, variable in size and habit,<br />

caespitose, creeping, erect to pendent, occasionally<br />

with pseudobulbs. LEAVES distichously arranged<br />

along the stem, occasionally one apical leaf or several<br />

apical leaves distributed throughout the stem or<br />

aggregate at the apex <strong>of</strong> the stem or pseudobulb,<br />

conduplicate, subcoriaceous to fleshy, not petiolate.<br />

INFLORESCENCE apical, rarely lateral, a one- to manyflowered<br />

raceme, spike, panicle or umbel. FLOWERS<br />

resupinate or not, small to large and showy. SEPALS<br />

AND PETALS subsimilar, free, usually spreading; the<br />

lip simple or lobed, mostly united to the ventral<br />

portion <strong>of</strong> the column, usually with fleshy calli at the<br />

base. COLUMN mostly totally fused with the lip, but<br />

sometimes totally free or half fused with the lip just<br />

to the apex, <strong>of</strong>ten provided with a hooded clinandrium.<br />

POLLINIA two, four or rarely eight, waxy.<br />

Epidendrum is one <strong>of</strong> the largest genera <strong>of</strong> Neotropical<br />

<strong>orchid</strong>s with more than 1500 species. <strong>The</strong> genus<br />

is widely distributed throughout the Neotropics, from<br />

North Carolina to northern Argentina including the<br />

Antilles.<br />

1. EPIDENDRUM COCOENSE HÁGSATER, Icon. Orchid.<br />

(Mexico) 3: pl. 325. 1999. Type: COSTA RICA, PUN-<br />

TARENAS, <strong>Cocos</strong> <strong>Island</strong>, along brook flowing into<br />

Wafer Bay, on trees, 18 April 1930, H.K. Svenson<br />

333 (holotype: AMES; isotypes: AMES, BKL, F).<br />

(Figs 3C, 4).<br />

Description: PLANT a large epiphyte, pendent, with<br />

terete, leafy stem up to 2.5 m long. Roots thick, up<br />

to 1–2 mm in diameter, white to greenish, with green<br />

tips, produced basally. STEM terete, branching, canelike,<br />

with lateral, secondary stems shorter than the<br />

main stems, each stems ending in apical inflorescences.<br />

LEAF SHEATHS tubular, enclosing the stem,<br />

laterally flattened, to 3 cm long. LEAVES many (to 30),<br />

distichous, distributed throughout the stem, laterally<br />

twisted, lanceolate, elliptic–oblong, acute, with a<br />

bilobed apex, conduplicate, subcoriaceous, articulate<br />

with the sheath involving the stem, to approximately<br />

5–14 ¥ 1.5–2.7 cm. INFLORESCENCE apical, racemose,<br />

distichous, two- to four-flowered, shorter than the<br />

leaves, flowering once, to 6 cm long, covered by<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


ORCHID FLORA OF COCOS ISLAND 29<br />

Figure 4. Epidendrum cocoense Hágsater. A, habit. B, segment <strong>of</strong> the inflorescence. C, flower. D, dissected perianth. E,<br />

column and lip, lateral view. F, column, lateral and front views. Drawn by D. Bogarín from Bogarín 2767 (JBL-Spirit).<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


30 D. BOGARÍN ET AL.<br />

imbricate sheaths, foliaceous, becoming papyraceous<br />

with age. PEDICEL inconspicuous, < 0.5 mm long,<br />

<strong>flora</strong>l bracts widely ovate, longer than the ovary,<br />

scarious, conduplicate, acute to subacute, 2.5–<br />

3.3 ¥ 1.5–2.0 cm. OVARY cylindrical, to 2 ¥ 3 mm,<br />

scurfy. FLOWERS white–cream, approximately<br />

1.7 cm in length. DORSAL SEPAL subequal to the<br />

lateral sepals, rectangular–ovate to elliptic, acute,<br />

12 ¥ 5 mm. LATERAL SEPALS ovate, elliptic, acute,<br />

slightly mucronate, 10 ¥ 5 mm. PETALS rectangular–<br />

oblong, subacute, margins revolute, 10 ¥ 2.5 mm. LIP<br />

triangular to slightly cordate, with basal rounded<br />

lobes covering the column apex, subacute, callus<br />

basal, rectangular with erose apex touching the<br />

column apex and a central keel running to the lip<br />

apex, adnate to the column, 1.0 ¥ 0.7 cm. COLUMN<br />

short, straight, slightly alate, with sinuose thick<br />

wings, widely dilated towards the apical half in<br />

lateral position, with an minutely acute, triangular<br />

tooth on each side, 5 ¥ 4 mm, anther apical, stigma<br />

ventral. CLINANDRIUM reduced, erose, sinuose.<br />

NECTARY prominent, penetrating the ovary. POLLINIA<br />

four, ovoid in pairs, with caudicles. ANTHER CAP cucullate,<br />

four-celled. CAPSULE pyriform, 3.0 ¥ 1.5 cm.<br />

Other vouchers examined: COSTA RICA. Puntarenas:<br />

Puntarenas, Isla del Coco, camino al Cerro Iglesias,<br />

cerca de Llanos Palo de Hierro, 310 m, 5°31′55.3N,<br />

87°03′47.8″W, bosque pluvial premontano, transición<br />

a basal, epífitas en Sacoglottis holdridgei, 15 abril<br />

2006, D. Bogarín 2750, G. Blanco, G. Gigot, V. Savolainen<br />

& J. Warner (JBL-Spirit). Puntarenas: Puntarenas,<br />

Isla del Coco, orillas del Río Genio, camino a<br />

la represa, 30 m, 5°32′33.08″N, 87°03′10.3″W, bosque<br />

pluvial premontano, transición a basal, epífita en<br />

Sacoglottis holdridgei, 15 abril 2006, D. Bogarín<br />

2732, G. Blanco, G.Gigot, V. Savolainen & J. Warner<br />

(JBL-Spirit). Puntarenas: Puntarenas, Isla del Coco,<br />

orillas del Río Genio, cerca de la casa de máquinas<br />

de la planta hidroeléctrica, 15 m, 5°32′60.08″N,<br />

87°03′41.3″W, bosque pluvial premontano, transición<br />

a basal, epífitas, 15 abril 2006, D. Bogarín 2767, G.<br />

Blanco, G. Gigot, V. Savolainen & J. Warner (JBL-<br />

Spirit). Puntarenas: Puntarenas, Isla del Coco, orillas<br />

del Río Genio, 50 m a la izquierda de la Casa de<br />

Máquinas, 30 m, 5°32′31.4″N, 87°03′16.7″W, bosque<br />

pluvial premontano, transición a basal, 18 abril 2007,<br />

D. Bogarín 3704, G.Gigot & J.D. Zuñiga (JBL-Spirit).<br />

Wafer Bay, 14 August 1973, R.L. Dressler 4471 (CR).<br />

Isla del Coco, bosque de Wafer, 20 m, febrero 1970,<br />

L.D. Gómez 3270 (CR). Isla del Coco, Bahía wafer, c.<br />

nivel del mar, 31 Julio 1981, J. Gómez-Laurito 6914<br />

(CR). Wafer Bay, June 28, 1932, J.T. Howell 10190<br />

(CAS). <strong>Cocos</strong> <strong>Island</strong>, December 5, 1959, W. Klawe<br />

1519 (US). Lépiz 339 (INB). Sendero Cerro Iglesias,<br />

29 June 1997, A. Rojas 3691 (CR, INB). Bahía Wafer,<br />

21 enero 1980, P. Sánchez-Vindas 10 (CR). Isla del<br />

Coco, 6–8 junio 1989, R. Soto s.n. (USJ). <strong>Cocos</strong> <strong>Island</strong>,<br />

along brook flowing into Wafer Bay, on trees, 18 April<br />

1930, H. K. Svenson 333 (AMES, BKL, F). Isla del<br />

Coco, 5°32′N, 87°03′W, November 2002, J. Trusty 101,<br />

167, 176, 184, 194, 195, 363, 387, 461, 483, 494, 496,<br />

501, 510, 511, 512, 520, 544, 548, 560 and 561 (CR,<br />

FTG). Isla del Coco, marzo 1940, J. Valerio 1108 (CR).<br />

Distribution: known only from <strong>Cocos</strong> <strong>Island</strong> in Costa<br />

Rica. Most plants are found between sea level and<br />

250 m elevation, but some can reach 575 m elevation<br />

on Cerro Iglesias.<br />

Etymology: from the Spanish word ‘coco’, coconut the<br />

name <strong>of</strong> the island to which this species is endemic.<br />

Habitat and ecology: A common endemic epiphyte in<br />

premontane rainforest, basal belt transition. Plants<br />

are easily found on the common <strong>Cocos</strong> <strong>Island</strong> endemic<br />

Sacoglottis holdridgei. <strong>The</strong>y grow hanging, mostly<br />

(but not strictly) on the main tree trunks at 2 m or<br />

less above the ground in shade understorey conditions.<br />

However, other individuals can be found in<br />

exposed humid conditions around the Genio River,<br />

Wafer and Chatham Bays. <strong>The</strong>y are recorded across<br />

the island, but in some areas with scattered Clusia<br />

rosea trees and rocky cliffs such as Punta Presidio<br />

there were no plants.<br />

Phenology: Flowers are produced from December to<br />

August.<br />

Discussion: Among the <strong>orchid</strong>s <strong>of</strong> the island, E.<br />

cocoense can be distinguished by the large pendent<br />

plants up to 2.5 m long. It can be distinguished from<br />

E. insulanum and E. jimenezii by the larger main<br />

stems, its conspicuous broad <strong>flora</strong>l bracts to 1.3–<br />

2.0 cm wide (vs. less than 0.5–0.6 cm wide in E.<br />

insulanum and E. jimenezii) and the scurfy ovary<br />

5 mm in diameter (vs. a glabrescent ovary < 2mmin<br />

diameter). <strong>The</strong> leaves are wider and longer, up to<br />

2.7 cm wide and 12.5 cm long (vs. < 1.5 cm wide and<br />

8 cm long in the other Epidendrum spp.). Also, it<br />

differs from E. insulanum in the inflorescence having<br />

three or four white flowers, rarely two (vs. one yellow<br />

flower), the column with two apical teeth in each side<br />

(vs. erose column apex) and the lateral lobes <strong>of</strong> lip<br />

covering the column apex in lateral position (vs.<br />

lateral lobes <strong>of</strong> lip not covering the column apex). It<br />

differs from E. jimenezii by the less conspicuous teeth<br />

at the apex <strong>of</strong> the column, the subacute lip apex and<br />

the wider <strong>flora</strong>l bracts completely covering the ovary.<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


Conservation status: <strong>The</strong> first IUCN Red List Assessment<br />

<strong>of</strong> E. cocoense by Trusty (2004) estimated an<br />

extent <strong>of</strong> occurrence (EOO) <strong>of</strong> 22.3 km 2 . Based on<br />

fieldwork collections during a trip in 2006, the EOO<br />

was calculated as 4.5 km 2 and the area <strong>of</strong> occupancy<br />

(AOO) in 7 km 2 . However, as E. cocoense is a common<br />

epiphyte in the island, its distribution in the whole<br />

territory is expected, with the exception <strong>of</strong> few disturbed<br />

areas and cliffs along the coast. If threats are<br />

identified in the future, then a rating <strong>of</strong> Vulnerable<br />

(VU) under criterion D2 may be appropriate. In 2004,<br />

Trusty suggested classifying this species either as<br />

Critically Endangered (CR) or as VU because <strong>of</strong> criterion<br />

B and D. <strong>The</strong> current conservation status by<br />

IUCN is Near Threatened (NT).<br />

2. EPIDENDRUM INSULANUM SCHLTR., Beih. Bot.<br />

Centralbl., Abt. 36(2): 404. 1918. Type: Costa Rica.<br />

Cordon littoral à Wafer Bay, <strong>Cocos</strong> <strong>Island</strong> (Pacific<br />

Ocean), January 1902, H. Pittier (n. 16350 Herb.<br />

Nacion. Costa Rica). Lectotype GH-3580 (=AMES-<br />

73449), designated by Trusty & Blanco (Epidendrum<br />

27: 12–13. 2005) Isosyntypes B (destroyed, drawing in<br />

AMES, AMES-70447), CR (two sheets), AMES [two<br />

sheets, GH-3581 (=AMES-70445), GH-3579 (=AMES-<br />

70446). (Figs 3D, 5).<br />

Description: PLANT epiphytic, pendent, erect or<br />

suberect, with terete, leafy stem to 30 cm long. Roots<br />

thin, < 1.5 mm in diameter, white to greenish, with<br />

green tips, produced basally. STEM terete, branching,<br />

cane-like, with lateral, secondary stems shorter than<br />

the main stems, each stems ending in apical inflorescences.<br />

LEAF SHEATHS tubular, enclosing the stem,<br />

laterally flattened, to 1 cm long. LEAVES many (to 18),<br />

distichous, distributed throughout the stem, laterally<br />

twisted, lanceolate, elliptic–oblong, acute, with a<br />

bilobed apex, conduplicate, coriaceous, articulate with<br />

the sheath involving the stem, greenish to yellowish,<br />

to c. 5–6 ¥ 2.5–1.3 cm. INFLORESCENCE apical, racemose,<br />

distichous, one-flowered, shorter than the<br />

leaves, flowering once, to 5 cm long, covered by<br />

imbricate sheaths, foliaceous, papyraceous with age;<br />

pedicel inconspicuous, < 0.5 mm long. FLORAL BRACTS<br />

tubular, slightly larger than the ovary, scarious, acute<br />

to subacute, to 6 mm long. OVARY cylindric, to 5 cm<br />

long. FLOWERS yellowish to lemon, approximately<br />

6 mm in length. DORSAL SEPAL subequal to the lateral<br />

sepals, ovate to elliptic, subacute, 7.5 ¥ 2.3 mm.<br />

LATERAL SEPALS ovate to elliptic, acute, 7.5 ¥ 2.5 mm.<br />

PETALS anguste–oblong to lanceolate, acute, margins<br />

slightly revolute, 7.0 ¥ 1.2 mm. LIP subcordate, with<br />

short basal rounded lobes not covering the column<br />

apex and produced just after the apex <strong>of</strong> the column,<br />

acute, callus basal, triangular with touching the<br />

column apex and a central keel running to the lip<br />

apex, adnate to the column, 6.0 ¥ 2.5 mm. COLUMN<br />

ORCHID FLORA OF COCOS ISLAND 31<br />

very short, straight, almost tubular, without teeth,<br />

5.0 ¥ 3.3 mm, anther apical, stigma ventral. CLINAN-<br />

DRIUM reduced, erose, sinuose. NECTARY prominent,<br />

penetrating the ovary. POLLINIA four, ovoid in pairs,<br />

with caudicles. ANTHER CAP cucullate, four-celled.<br />

Other vouchers examined: COSTA RICA. Puntarenas:<br />

Puntarenas, Isla del Coco, Llanos Palo de Hierro,<br />

310 m, 5°31′55.3″N, 87°03′47.8″W, bosque pluvial premontano,<br />

transición a basal, epífitas en Sacoglottis<br />

holdridgei, 16 abril 2006, D. Bogarín 2772, G. Blanco,<br />

G.Gigot, V. Savolainen & J. Warner (JBL-Spirit). Puntarenas:<br />

Puntarenas, Isla del Coco, camino al Cerro<br />

Iglesias, cerca de Llanos Palo de Hierro, 310 m,<br />

5°31′55.3″N, 87°03′47.8″W, bosque pluvial premontano,<br />

transición a basal, epífitas en Sacoglottis holdridgei,<br />

15 abril 2006, D. Bogarín 2762, G. Blanco,<br />

G.Gigot, V. Savolainen & J. Warner (JBL-Spirit). Puntarenas:<br />

Puntarenas, Isla del Coco, orillas del Río<br />

Genio, 50 m a la izquierda de la Casa de Máquinas,<br />

30 m, 5°32′31.4″N, 87°03′16.7″W, bosque pluvial premontano,<br />

transición a basal, 18 abril 2007, D. Bogarín<br />

3707, G. Gigot & J.D. Zuñiga (JBL-Spirit). Same<br />

locality: D. Bogarín 3705, 3708, 3702, 3706. (JBL-<br />

Spirit). <strong>Cocos</strong> <strong>Island</strong>, 28 march 1891, A. Agassiz s.n.<br />

(GH) near Wafer Bay, 14 august 1973, R.L. Dressler<br />

4468 (CR, PMA). 9–11 April 1979, R.B Foster 4117 (F,<br />

US). Isla del Coco, 31 July 1981, J. Gómez-Laurito<br />

6924 (CR). Río Genio, Estación Wafer, 1–10 m, 14<br />

abril 1994, E. Lépiz 358 (INB). Cerro Iglesias, 200–<br />

250 m, 22 Junio 1997, A. Rojas 3660 (F, INB). Wafer<br />

Bay, 18 April 1930, H. Svenson 335 (AMES, BKL, F,<br />

K, LE, S, UC). Isla del Coco, 5°31′N, 87°03′W,<br />

J. Trusty 53, 158, 218 and 500 (CR, USJ).<br />

Distribution: known only from <strong>Cocos</strong> <strong>Island</strong>, Costa<br />

Rica.<br />

Etymology: From the Latin insula, ‘island’, in allusion<br />

to <strong>Cocos</strong> <strong>Island</strong>, to which this species is endemic.<br />

Phenology: Apparently they flower throughout the<br />

year. However, plants collected and cultivated for this<br />

study flowered from March to July.<br />

Habitat and ecology: A common endemic epiphyte<br />

in premontane rainforest, basal belt transition. <strong>The</strong><br />

habitat is mostly the same as for E. cocoense and<br />

sometimes they grow intermixed. <strong>The</strong>y are pendent or<br />

suberect, growing on exposed or shady conditions in<br />

both shrubs and trees. Most <strong>of</strong> the plants are found<br />

between sea level and 300 m <strong>of</strong> elevation; however,<br />

they can reach 575 m on Cerro Iglesias. Pollinators<br />

are unknown. <strong>The</strong> three Epidendrum spp. have<br />

similar <strong>flora</strong>l architecture. As in E. cocoense and E.<br />

jimenezii, plants could be pollinated by Lepidoptera.<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


32 D. BOGARÍN ET AL.<br />

Figure 5. Epidendrum insulanum Schltr. A, habit. B, segment <strong>of</strong> the inflorescence. C, flower. D, dissected perianth. E,<br />

column and lip, lateral view. F, column, front view. Drawn by D. Bogarín from Bogarín 2772 (JBL-Spirit).<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


Discussion: Epidendrum insulanum is easily distinguished<br />

by the inflorescences having only one, green–<br />

yellow flower (vs. whitish to white, and two or four in<br />

E. jimenezii and E. cocoense, respectively). <strong>The</strong> apex<br />

<strong>of</strong> column is erose, lacking the two peaks present in<br />

the other species. <strong>The</strong> lateral lobes <strong>of</strong> lip do not cover<br />

the apex <strong>of</strong> column in lateral view. <strong>The</strong> lip is smaller,<br />

up to 6 mm long. Sepals and petals are smaller,<br />

< 1 cm in length. Vegetatively, plants are smaller than<br />

those <strong>of</strong> E. cocoense, reaching up to 30 cm long with<br />

leaves < 1.5 cm wide. Some larger plants can be confused<br />

with E. jimenezii because <strong>of</strong> the similar size.<br />

Conservation status: In 2004, Trusty suggested classifying<br />

this species either as Critically Endangered<br />

CR or as VU because <strong>of</strong> criterion B and D. Based on<br />

herbarium specimens and field collections, she estimated<br />

an EOO <strong>of</strong> 18.2 km 2 . Based on field collections<br />

performed in 2006, the EOO was calculated as 3 km 2<br />

and the AOO as 7 km 2 . Although restricted in range<br />

and variation in EOO and AOO values, based on field<br />

observations it is highly probable that plants <strong>of</strong> this<br />

species are found on the whole island. <strong>The</strong>y are not<br />

restricted to a particular place. <strong>The</strong> current conservation<br />

status by IUCN is Near Threatened (NT).<br />

3. EPIDENDRUM JIMENEZII HÁGSATER, Icon. Orchid.<br />

(Mexico) 3: pl. 341. 1999. Type: Costa Rica. <strong>Cocos</strong><br />

<strong>Island</strong>, Wafer Bay, Arroyo del Genio, 13 April 1965, A.<br />

Jiménez M. 3178 (holotype: F isotypes: CR, MO, SEL,<br />

U) (Figs 3E, 6).<br />

Description: PLANT epiphytic, pendent, with terete,<br />

leafy stem to 35 cm long. ROOTS thin, < 1.5 mm in<br />

diameter, white to greenish, with green tips, produced<br />

basally. STEM terete, branching, cane-like, with<br />

lateral, secondary stems shorter than the main stems,<br />

each stem ending in apical inflorescences. LEAF<br />

SHEATHS tubular, enclosing the stem, laterally flattened,<br />

to 2 cm long. LEAVES many (to 20), distichous,<br />

distributed throughout the stem, laterally twisted,<br />

lanceolate, elliptic–oblong, acute, with a bilobed apex,<br />

conduplicate, coriaceous to subcoriaceous, articulate<br />

with the sheath involving the stem, to approximately<br />

6–8 ¥ 1.0–1.3 cm. INFLORESCENCE apical, racemose,<br />

distichous, two-flowered, shorter than the leaves,<br />

flowering once, to 3.1 cm long, covered by imbricate<br />

sheaths, foliaceous, becoming papyraceous with<br />

age. PEDICEL inconspicuous, < 0.5 mm long. FLORAL<br />

BRACTS tubular, as long as the ovary, scarious, acute<br />

to subacute, 0.5–0.6 cm long. OVARY cylindric, to<br />

1.5 cm long. FLOWERS whitish, about 1.3 cm in length.<br />

DORSAL SEPAL subequal to the lateral sepals, ovate to<br />

elliptic, acute, 9.5 ¥ 3.0 mm. LATERAL SEPALS ovate to<br />

elliptic, acute, 9.5 ¥ 3.0 mm. PETALS rectangular–<br />

oblong to lanceolate, acute to subacute, margins revolute,<br />

8 ¥ 2 mm. LIP cordate, with basal rounded lobes<br />

ORCHID FLORA OF COCOS ISLAND 33<br />

covering the column apex, acute, callus basal, rectangular<br />

with erose apex touching the column apex and<br />

a central keel running to the lip apex, adnate to the<br />

column, 7 ¥ 4 mm. COLUMN short, straight, slightly<br />

alate, with sinuose wings, widely dilated towards<br />

the apical half in lateral position, with an acute,<br />

triangular tooth on each side, 5.0 ¥ 2.2 mm, anther<br />

apical, stigma ventral. CLINANDRIUM reduced,<br />

sinuose. NECTARY prominent, penetrating the ovary.<br />

POLLINIA four, ovoid in pairs, with caudicles. ANTHER<br />

CAP cucullate, four-celled.<br />

Other vouchers examined: COSTA RICA. Puntarenas:<br />

Puntarenas, Isla del Coco, orillas del Río Genio,<br />

camino a la represa, 30 m, 5°32′33.08N, 87°03′10.3″W,<br />

bosque pluvial premontano, transición a basal,<br />

epífita, 13 abril 2006, D. Bogarín 2735, G. Blanco,<br />

G.Gigot, V. Savolainen & J. Warner (JBL-Spirit).<br />

<strong>Cocos</strong> <strong>Island</strong>, Wafer Bay, Arroyo del Genio, 13 April<br />

1965, A. Jiménez M. 3178 (CR); Wafer Bay, R.L.<br />

Dressler 4468 (FLAS); <strong>Cocos</strong> <strong>Island</strong>, R. Foster 4114<br />

(US).<br />

Distribution: Known only from <strong>Cocos</strong> <strong>Island</strong>, Costa<br />

Rica. It is restricted to the Genio River and Wafer Bay<br />

areas.<br />

Eponymy: Named in honour <strong>of</strong> Ing. Alfonso Jiménez<br />

Muñoz, ex-pr<strong>of</strong>esor <strong>of</strong> Agronomy and Biology at Universidad<br />

Nacional de Costa Rica and ex-director <strong>of</strong><br />

the Museo Nacional de Costa Rica, who collected the<br />

type specimen.<br />

Phenology: December to January.<br />

Habitat and ecology: a very rare endemic epiphytic<br />

<strong>orchid</strong> in premontane rainforest, basal belt transition.<br />

Only five specimens are known from the island and<br />

their ecology is poorly understood. As noted for the<br />

other Epidendrum spp., the <strong>flora</strong>l architecture is<br />

similar and their pollinators could share the same<br />

characteristics. <strong>The</strong> only plant found in this study<br />

was growing along the path to the Genio River waterfall,<br />

in shady humid conditions.<br />

Discussion: Plants <strong>of</strong> this species are distinguished by<br />

the inflorescence producing two whitish flowers, the<br />

cordate, acute lip and the column with two acute<br />

teeth at apex. Compared with E. cocoense, it can be<br />

distinguished by its smaller habit, up to 35 cm long,<br />

the leaves 1.0–1.2 wide, the less conspicuous tubular<br />

<strong>flora</strong>l bracts 0.5–0.6 cm wide and as long as the glabrescent<br />

ovary. Vegetatively it is similar to some<br />

larger plants <strong>of</strong> E. jimenezii. Large plants <strong>of</strong> E. insulanum<br />

are <strong>of</strong>ten indistinguishable from those <strong>of</strong><br />

E. jimenezii. However, plants <strong>of</strong> E. jimenezii can be<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


34 D. BOGARÍN ET AL.<br />

Figure 6. Epidendrum jimenezii Hágsater. A, habit. B, segment <strong>of</strong> the inflorescence. C, flower. D, dissected perianth. E,<br />

column and lip, lateral view. F, column, lateral and front views. Drawn by D. Bogarín from Bogarín 2735 (JBL-Spirit).<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


distinguished by the inflorescences having two<br />

whitish flowers, > 1 cm in length, the column apex<br />

with two teeth in each side and the lateral lobes <strong>of</strong> lip<br />

covering the column apex.<br />

<strong>The</strong> <strong>flora</strong>l features <strong>of</strong> the three Epidendrum spp.<br />

are similar. <strong>The</strong> only specimen collected for this study<br />

seems to be intermediate between E. cocoense and<br />

E. insulanum. Basically, the lip shape is similar to<br />

E. cocoense, with the same callus structure, but<br />

having rounded basal lobes and acute apex as in<br />

E. insulanum. <strong>The</strong> sepals and petals are as long as<br />

those <strong>of</strong> E. cocoense, but the shape is more similar to<br />

E. insulanum. Plant size is intermediate between the<br />

two species. <strong>The</strong> column shape also shares intermediate<br />

characteristics. <strong>The</strong> column is oblong as in<br />

E. insulanum but with two teeth at apex as in<br />

E. cocoense.<br />

Trusty (2006) was unable to distinguish this species<br />

from E. insulanum. <strong>The</strong>y suggested the possibility<br />

that this species could be a hybrid between E.<br />

cocoense and E. insulanum and observations <strong>of</strong> <strong>flora</strong>l<br />

morphology support their hypothesis. However, more<br />

evidence is needed to complement the morphological<br />

data, especially at population genetics and reproductive<br />

biology levels.<br />

Conservation status: Trusty (2004) suggested classifying<br />

this species as CR because <strong>of</strong> criterion B and D.<br />

<strong>The</strong> estimation <strong>of</strong> the EOO is < 1km 2 . This species is<br />

very rare and restricted to the Wafer Bay areas<br />

around the Genio River where it is known from less<br />

than five collections. This species has not been evaluated<br />

yet under IUCN criteria; however, it should be<br />

classified as CR as suggested by Trusty (2004).<br />

<strong>The</strong> endemic Epidendrum species: <strong>The</strong> three Epidendrum<br />

spp. are similar in their vegetative morphology.<br />

All the species have a pendent branching habit. <strong>The</strong><br />

inflorescence is terminal and new branches are produced<br />

beneath the apex after they have flowered. <strong>The</strong><br />

new branches flower again. Flowers are superficially<br />

similar. <strong>The</strong> lip is cordate or subcordate with the<br />

callus always Y-shaped. <strong>The</strong> column has a reduced<br />

clinandrium and shows little variation between<br />

species. <strong>The</strong> perianth segments show some variation<br />

in size and their shape varies from oblong to<br />

ovate–elliptic.<br />

Relationships with other members <strong>of</strong> the genus<br />

Epidendrum: According to Hágsater (2007), they are<br />

members <strong>of</strong> the Epidendrum ramosum Jacq. group,<br />

which is characterized by the monopodial branching<br />

stems, the spike-like distichous inflorescence and the<br />

lip with a single callus and the flexicaule subgroup,<br />

ORCHID FLORA OF COCOS ISLAND 35<br />

which has a straggling habit, the main stem not very<br />

evident and the dorsal keel present in the sepals,<br />

which is generally prominent.<br />

Epidendrum insulanum is closely related to E.<br />

flexicaule Schltr., E. stevensii Hágsater, E. modestiflorum<br />

Schltr and E. veraguasense Hágsater (Hágsater,<br />

2007), but they can be distinguished: E. flexicaule has<br />

longer leaves (up to 8.2 cm long) and two or three<br />

larger flowers, E. stevensii has two or three smaller<br />

flowers, E. modestiflorum shorter wider leaves<br />

(5 ¥ 15 cm) and two or three flowers and E. veraguasense<br />

has two or three white, larger flowers. Epidendrum<br />

jimenezii is perhaps closely allied to E.<br />

insulanum and E. cocense. Being sympatric and<br />

having similar plant size, E. jimenezii and E. insulanum<br />

are sometimes confused.<br />

Epidendrum cocoense is vegetatively similar to E.<br />

rafael-lucasii Hágsater, E. santaclarense Ames and E.<br />

acunae Dressler. <strong>The</strong>y are found in Central America<br />

and Panama, with the exception <strong>of</strong> the Costa Rican<br />

endemic E. rafael-lucasii. According to Hágsater<br />

(1999a), E. rafael-lucasii has larger yellow flowers (vs.<br />

white), sepals 16–20 mm long (vs. 12 mm in E.<br />

cocoense) and the column is blunt at the apex (vs.<br />

with two teeth at each side <strong>of</strong> apex). Epidendrum<br />

santaclarense has deep green flowers (vs. white) and<br />

E. acunae has shorter stems and very short flowering<br />

stems.<br />

Pollination <strong>of</strong> endemic species <strong>of</strong> Epidendrum: <strong>The</strong><br />

pollination biology is unknown. Epidendrum spp.<br />

have a deep tube inside the ovary, sometimes with<br />

free nectar. <strong>The</strong> lip and column made up a simple<br />

horizontal tube, with the free part <strong>of</strong> the lip used as<br />

an appendage for pollinator attraction. Some species<br />

<strong>of</strong> Lepidoptera, mainly moths, are important pollinators<br />

<strong>of</strong> members <strong>of</strong> subtribe Laeliinae Benth., including<br />

Epidendrum (van der Pijl & Dodson, 1966). <strong>The</strong><br />

white to whitish flowers <strong>of</strong> E. cocoense produce a<br />

perceptible sweet smell and they could attract some<br />

species <strong>of</strong> moths or butterflies. Nearly 100 species<br />

<strong>of</strong> Lepidoptera are recorded from <strong>Cocos</strong> <strong>Island</strong> and<br />

moths are the largest group (Brown, Donahue &<br />

Miller, 1991; Brown & Miller, 1999). As there are no<br />

bees in the island, it is highly probable that the<br />

Epidendrum spp. are pollinated by Lepidoptera. Selfpollination<br />

was not observed and plants in cultivation<br />

flowered without producing fruits.<br />

3. ORNITHIDIUM SALISB. EX R.BR.<br />

ORNITHIDIUM SALISB. EX R.BR., Hort. Kew. ed. 2, 5:<br />

210. 1813. Type species: Epidendrum coccineum Jacq.,<br />

Enum. Syst. Pl. 29. 1760. = Ornithidium coccineum<br />

(Jacq.) Salisb., Trans. Hort. Soc. London 1: 293. 1812.<br />

EPIPHYTIC, sympodial (caespitose to rhizomatous)<br />

or monopodial plants; rarely with dimorphic growth.<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


36 D. BOGARÍN ET AL.<br />

<strong>The</strong> thick roots with an orangish coloration. STEMS<br />

cane-like with distichous leaves or with pseudobulbs<br />

bearing basal and apical leaves. <strong>The</strong> stems and leaves<br />

<strong>of</strong> most species have an olive green coloration. LEAVES<br />

conduplicate, coriaceous to subcoriaceous. INFLORES-<br />

CENCES usually fascicled, pedicel and ovary longer<br />

than the <strong>flora</strong>l bract. FLOWERS usually small, fleshy,<br />

campanulate or subglobose, the perianth lacks fibres;<br />

many species have yellow, orange, red, tan, green or<br />

pink flowers. SEPALS AND PETALS subsimilar, spreading.<br />

LIP simple or three- or four-lobed, articulated<br />

with or adnate to the column, usually provided at the<br />

base with a fleshy callus and <strong>of</strong>ten producing nectar<br />

at the base <strong>of</strong> the labellum. COLUMN erect, terete.<br />

CAPSULES have apical dehiscence.<br />

Ornithidium is a widely distributed genus throughout<br />

the Neotropics with approximately 60 species.<br />

1. ORNITHIDIUM ADENDROBIUM (RCHB.F.)<br />

M.A.BLANCO &OJEDA, Lankesteriana 7: 532. 2007.<br />

Ponera adendrobium Rchb.f., Flora 48: 278. 1865.<br />

Pleuranthium adendrobium (Rchb.f.) Benth. &<br />

Hook.f. ex B.D.Jacks., Index Kew. 2: 562. 1894. Neourbania<br />

adendrobium (Rchb.f.) Fawc. & Rendle, J.<br />

Bot. 47: 125 1909. Type: CUBA. 1860–1864, C. H.<br />

Wright 1697 (holotype: W; isosintypes: AMES, BM, K,<br />

MO). Camaridium parviflorum Fawc. in I. Urban,<br />

Symb. Antill. 1: 472. 1900. (Figs 3F, 7).<br />

Description: PLANT epiphytic, erect or suberect, with<br />

terete, leafy stem, to 60 cm long. ROOTS flexuous,<br />

less < 1.5 mm in diameter, orangish to whitish. STEM<br />

without pseudobulbs, terete, leafy, covered by<br />

addressed leaf sheaths, scarious at base, foliaceous<br />

towards the apex, producing secondary branches,<br />

with adventitious roots produced at bases <strong>of</strong> leaf<br />

nodes. LEAVES many (to 25), distichous, narrowly<br />

oblong to anguste–elliptic, acute, conduplicate, fleshy,<br />

flexible, articulate with the sheath involving the<br />

stem, to c. 11.0 ¥ 1.6 cm. INFLORESCENCE a singleflowered<br />

raceme usually larger than the leaves, produced<br />

along the stem, in the axils <strong>of</strong> the leaves, c.<br />

2 cm long. PEDICEL inconspicuous, < 0.5 mm long.<br />

FLORAL BRACTS ovate, foliaceous. OVARY cylindric,<br />

< 4 mm long, glabrous. FLOWERS small, inconspicuous,<br />

c. 6 mm in length, yellowish to cream–yellow<br />

or greenish. DORSAL SEPAL subequal to the lateral<br />

sepals, oblong–elliptic, acute or slightly acuminate,<br />

6 ¥ 2 mm. LATERAL SEPALS oblong–elliptic, acute,<br />

subfalcate, 6 ¥ 2 mm. PETALS ovate–lanceolate, acute,<br />

parallel to the column, 5.0 ¥ 1.7 mm. LIP ovate, lobed,<br />

sinuose, retuse, margin erose, callous laminar in the<br />

centre, with two parallel keels running to the apex,<br />

attached to the column foot. COLUMN short, terete,<br />

slightly arcuate, to 3 mm long; stigma ventral, widely<br />

obovate, anther terminal. POLLINIA two. ANTHER CAP,<br />

cucullate. CAPSULE pyriform to 1.7 cm long.<br />

Other vouchers examined: COSTA RICA. Puntarenas:<br />

Puntarenas, Isla del Coco, camino al Cerro Iglesias,<br />

cerca de Llanos Palo de Hierro, 310 m, 5°31′55.3″N,<br />

87°03′47.8″W, bosque pluvial premontano, transición<br />

a basal, epífitas en Sacoglottis holdridgei, 15 abril<br />

2006, D. Bogarín 2755, G. Blanco, G. Gigot, V. Savolainen<br />

& J. Warner (JBL-Spirit). Same locality, D.<br />

Bogarín 2759 (JBL-Spirit). Puntarenas, Isla del Coco,<br />

December 2003, J. Trusty 495 (JBL-Spirit). Same<br />

locality: J. Trusty 52 (JBL-Spirit). Isla del Coco: Wafer<br />

Bay, 14 august 1973, R.L. Dressler 4470 (CR). Isla del<br />

Coco, J. Trusty 52, 100, 157, 495, 515, 556, 564 (CR).<br />

december 2003, J. Trusty 52 and 495 (JBL-spirit). Isla<br />

del Coco, región de los Llanos y subiendo a Cerro<br />

Pelón, 5°32′15″N 87°03′29″W, 200–250 m, 29 junio<br />

1997, A. Rojas 3692 (INB). 6–8 junio 1989, R. Soto<br />

s.n. (USJ-49286).<br />

Distribution: This is a widespread species ranging<br />

from Mexico and the Antilles throughout Central and<br />

South America.<br />

Etymology: Made up by the Latin prefix ‘a’, ‘away<br />

from’ and the name <strong>of</strong> the genus Dendrobium. According<br />

to Reichenbach (1865), the specimen he studied<br />

‘has all characteristics <strong>of</strong> Dendrobium, therefore the<br />

name’.<br />

Phenology: <strong>The</strong> plants have been recorded in flower<br />

throughout the year, mostly between September and<br />

April<br />

Habitat and ecology: Plants are frequent in the<br />

island. <strong>The</strong>y are mostly found from 250 to 585 m <strong>of</strong><br />

elevation, up to the top <strong>of</strong> Cerro Iglesias. It is rarely<br />

found at sea level. Some plants are probably selfpollinated<br />

because they form capsules quickly and<br />

regularly in almost all flowers. In the field, small ants<br />

were observed around the flowers but without pollinia.<br />

Capsules are seen throughout the year.<br />

Discussion: Plants have distichously arranged leaves<br />

without pseudobulbs. Compared with Epidendrum<br />

spp., it is easily recognized by the lateral singleflowered<br />

inflorescences and the footed column. It is<br />

distinguished from C. micranthum by its larger habit,<br />

without pseudobulbs and the stem with distichously<br />

arranged leaves. <strong>The</strong> small flowers do not have the<br />

lip adnate to the column foot. This species was previously<br />

treated in Maxillaria; however, nomenclatural<br />

changes in Maxillariinae suggest its inclusion in<br />

Ornithidium (Blanco et al., 2007; Whitten et al.,<br />

2009).<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


ORCHID FLORA OF COCOS ISLAND 37<br />

Figure 7. Ornithidium adendrobium (Rchb.f.) M.A.Blanco & Ojeda. A, habit. B, flowers. C, dissected perianth. D, column<br />

and lip, lateral view. E, column, lateral and front view. Drawn by D. Bogarín from Bogarín 2755 (JBL-Spirit).<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


38 D. BOGARÍN ET AL.<br />

Conservation status: This species has a wide distribution<br />

range throughout the Neotropics. It is common<br />

in the island. It should not be regarded as endangered<br />

and may well fit the LC category for widespread and<br />

abundant taxa.<br />

ACKNOWLEDGEMENTS<br />

<strong>The</strong> present paper is part <strong>of</strong> the Project 814-BO-245,<br />

‘La <strong>flora</strong> de orquídeas del Parque Nacional Isla del<br />

Coco’ sponsored by the Vice Presidency <strong>of</strong> Research,<br />

University <strong>of</strong> Costa Rica and ‘Conservation and Monitoring<br />

<strong>of</strong> Meso-American Orchids’ supported by the<br />

Darwin Initiative, Department for Environment Food<br />

and Rural Affairs (DEFRA), UK. We thank the Costa<br />

Rican Ministry <strong>of</strong> Environment and Energy (MINAE),<br />

the <strong>National</strong> System <strong>of</strong> Conservation Areas (SINAC)<br />

throughout Área de Conservación Marina Isla del<br />

Coco (ACMIC) for issuing the Resolution Permit no.<br />

002-06 and the Scientific Passport no. 1281, under<br />

which the material was collected and studied, Mario<br />

Blanco and Robert L. Dressler for useful discussions<br />

and digital images <strong>of</strong> critical specimens, the curators<br />

<strong>of</strong> AMES, BM, CAS, CR, INB, JBL, K, PMA and USJ,<br />

the crew <strong>of</strong> Okeanos for providing transportation and<br />

facilities to the island, the park guards and personnel<br />

<strong>of</strong> <strong>Cocos</strong> <strong>Island</strong> <strong>National</strong> <strong>Park</strong>, especially Isaac Chinchilla,<br />

German Haugh and Lourdes Vargas for granting<br />

access to the island and help with fieldwork<br />

logistics, Guillermo Blanco who shared his knowledge<br />

and most <strong>of</strong> the fieldwork and Guillaume Gigot and<br />

Jose Daniel Zúñiga for helping during the fieldwork<br />

activities.<br />

REFERENCES<br />

Alfaro EJ. 2008. Ciclo diario y anual de variables troposféricas<br />

y oceánicas en la Isla del Coco, Costa Rica. Revista de<br />

Biología Tropical 56: 19–29.<br />

Bellon H, Sáenz R, Tournon J. 1984. K-Ar radiometric ages<br />

<strong>of</strong> lavas from <strong>Cocos</strong> <strong>Island</strong> (Eastern Pacific). Marine Geology<br />

54: M17–M23.<br />

Bentham G. 1844–1846. <strong>The</strong> botany <strong>of</strong> the voyage <strong>of</strong> H.M.S.<br />

Sulphur, under command <strong>of</strong> Captain Sir Edward Belcher,<br />

during the years 1836–42. Published under the authority <strong>of</strong><br />

the Lords Commissioners <strong>of</strong> the Admiralty. London, UK. Vol.<br />

12.<br />

Blanco MA, Carnevali G, Whitten WM, Singer RB,<br />

Koehler S, Williams NH, Ojeda I, Neubig KM, Endara<br />

L. 2007. Generic realignments in Maxillariinae (Orchidaceae).<br />

Lankesteriana 7: 515–537.<br />

Bogarín D, Pupulin F. 2007. Las orquídeas del Parque<br />

Nacional Barra Honda, Guanacaste, Costa Rica. Lankesteriana<br />

7: 446–449.<br />

Brown JW, Miller SE. 1999. A new species <strong>of</strong> Coelostathma<br />

Clemens (Lepidoptera: Tortricidae) from <strong>Cocos</strong> <strong>Island</strong>, Costa<br />

Rica, with comments on the phylogenetic significance <strong>of</strong><br />

abdominal dorsal pits in Sparganothini. Proceedings <strong>of</strong> the<br />

Entomological Society <strong>of</strong> Washington 101: 701–707.<br />

Brown JW, Donahue JP, Miller SE. 1991. Two new species<br />

<strong>of</strong> geometrid moths (Lepidoptera: Geometridae: Ennominae)<br />

from <strong>Cocos</strong> <strong>Island</strong>, Costa Rica. Natural History Museum<br />

<strong>of</strong> Los Angeles County, Contributions in Science 423: 11–18.<br />

Cortés J. 2008. Historia de la investigación marina de la Isla<br />

del Coco, Costa Rica. Revista de Biología Tropical 56: 1–18.<br />

Dressler RL. 2003. Orchidaceae. In: Hammel BE, Grayum<br />

MH, Herrera C, Zamora N, eds. Manual de plantas de Costa<br />

Rica. Vol. 3. Monocotiledóneas (Orchidaceae–Zingiberaceae).<br />

Monographs in Systematic Botany from the Missouri Botanical<br />

Garden. St. Louis, MO: Missouri Botanical Garden<br />

Press, 93:1–595.<br />

Garay LA. 1967. Studies in American <strong>orchid</strong>s VI. Botanical<br />

Museum Leaflets 21: 249–264.<br />

Hágsater E. 1999a. Epidendrum cocoënse Hágsater. Sub. pl.<br />

325. In: Hágsater E, Sánchez L, García-Cruz J, eds. Icones<br />

Orchidacearum (Mexico) 3. <strong>The</strong> genus Epidendrum: part 2.<br />

A second century <strong>of</strong> new species in Epidendrum. Mexico:<br />

Herbario AMO, 301–400.<br />

Hágsater E. 1999b. Epidendrum jimenezii Hágsater. Sub. pl.<br />

341. In: Hágsater E, Sánchez L, García-Cruz J, eds. Icones<br />

Orchidacearum (Mexico) 3. <strong>The</strong> genus Epidendrum: part 2.<br />

A second century <strong>of</strong> new species in Epidendrum. Mexico:<br />

Herbario AMO, 301–400.<br />

Hágsater E. 2007. Epidendrum insulanum Hágsater. Sub.<br />

948. In: Hágsater E, Sánchez L, eds. Icones Orchidacearum<br />

(Mexico) 9. <strong>The</strong> genus Epidendrum: part 6. Species new and<br />

old in Epidendrum. Mexico: Herbario AMO, 901–1000.<br />

Hogue CL, Miller SE. 1981. Entom<strong>of</strong>auna <strong>of</strong> <strong>Cocos</strong> <strong>Island</strong>,<br />

Costa Rica. Atoll Research Bulletin 250: 1–29.<br />

Holdridge LR. 1967. Life zone ecology. San José: Tropical<br />

Science Center.<br />

Holdridge LR. 1987. Ecología basada en zonas de vida, 3rd<br />

edn. San José: IICA.<br />

Lizano OG. 2008. Dinámica de aguas alrededor de la Isla del<br />

Coco, Costa Rica. Revista de Biología Tropical 56: 31–48.<br />

van der Pijl L, Dodson CH. 1966. Orchid flowers: their<br />

pollination and evolution. Coral Gables: University <strong>of</strong><br />

Miami Press.<br />

Pittier HF. 1898. Apuntamientos preliminares sobre la Isla<br />

del <strong>Cocos</strong>, posesión costarricense en el Océano Pacifico.<br />

Revista del Colegio Superior de Señoritas 2: 2–11.<br />

Pupulin F. 1998. Orchid florula <strong>of</strong> Parque Nacional Manuel<br />

Antonio, Quepos, Costa Rica. Revista de Biología Tropical<br />

46: 961–10317.<br />

Pupulin F. 2007. <strong>Epidendra</strong>, the botanical databases <strong>of</strong><br />

Jardín Botánico Lankester at the University <strong>of</strong> Costa Rica.<br />

Lankesteriana 7: 178–180.<br />

Pupulin F. 2009. <strong>Epidendra</strong>, la base de datos en línea del<br />

Jardín Botánico Lankester de la Universidad de Costa Rica.<br />

Orquideología XXVI: 89–94.<br />

Pupulin F. 2010. Orchidaceae werckleanae: typification <strong>of</strong><br />

Costa Rican <strong>orchid</strong> species described from collections by K.<br />

Wercklé. Botanical Journal <strong>of</strong> the Linnean Society 163:<br />

111–154.<br />

Reichenbach HG. 1865. Vorstudien zu einer Orchidographie<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39


der Antillen, besonders britischen Antheils. Neuheiten und<br />

eingehende Critiken bisheriger Literatur I. Neuheiten aus<br />

Cuba von Wright. Flora 48: 273–280.<br />

Schlechter R. 1918. Kritische Aufzählung der bisher aus<br />

Zentralamerika bekanntgeworden Orchideen. Beihefte zum<br />

Botanischen Centralblatt, Kassel 36: 404.<br />

Singer RA. 2003. Orchid pollination: recent developments<br />

from Brazil. Lankesteriana 7: 111–114.<br />

Svenson HK. 1935. Plants <strong>of</strong> the Astor Expedition, 1930<br />

(Galápagos and <strong>Cocos</strong> <strong>Island</strong>s). American Journal <strong>of</strong> Botany<br />

22: 208–277.<br />

Tosi J. 1969. Mapa ecológico según la clasificación de zonas de<br />

vida del mundo de L. R. Holdridge. San José: Instituto<br />

Geográfico Nacional.<br />

Trusty J. 2004. Plant biogeography and conservation on a<br />

ORCHID FLORA OF COCOS ISLAND 39<br />

tropical island: Isla del Coco, Costa Rica. D. Phil. <strong>The</strong>sis,<br />

Florida International University.<br />

Trusty J, Blanco MA. 2005. Las orquídeas de la Isla del<br />

Coco. Epidendrum 27: 10–15.<br />

Trusty J, Blanco MA. 2006. Orchids <strong>of</strong> Isla del Coco: five<br />

species native to an island in Costa Rica. Orchids 75:<br />

826–831.<br />

Trusty J, Kesler HC, Haug Delgado G. 2006. Vascular <strong>flora</strong><br />

<strong>of</strong> Isla del Coco, Costa Rica. Proceedings <strong>of</strong> the California<br />

Academy Sciences 57: 247–355.<br />

Whitten WM, Blanco MA, Williams NH, Koehler S,<br />

Carnevali G, Singer RB, Endara L, Neubig KM. 2009.<br />

Molecular phylogenetics <strong>of</strong> Maxillaria and related genera<br />

(Orchidaceae: Cymbidiae) based on combined molecular<br />

data sets. American Journal <strong>of</strong> Botany 94: 1860–1889.<br />

© 2011 <strong>The</strong> Linnean Society <strong>of</strong> London, Botanical Journal <strong>of</strong> the Linnean Society, 2011, 166, 20–39

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